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The LIM and POU Homeobox Genes Ttx-3 and Unc-86 Act As Terminal

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The LIM and POU Homeobox Genes Ttx-3 and Unc-86 Act As Terminal © 2014. Published by The Company of Biologists Ltd | Development (2014) 141, 422-435 doi:10.1242/dev.099721 RESEARCH ARTICLE The LIM and POU homeobox genes ttx-3 and unc-86 act as terminal selectors in distinct cholinergic and serotonergic neuron types Feifan Zhang1, Abhishek Bhattacharya1, Jessica C. Nelson2, Namiko Abe1, Patricia Gordon1, Carla Lloret-Fernandez3, Miren Maicas3, Nuria Flames1,3, Richard S. Mann1, Daniel A. Colón-Ramos2 and Oliver Hobert1,4,* ABSTRACT factor initiates and maintains the terminal differentiation program of Transcription factors that drive neuron type-specific terminal dopaminergic neurons in the midbrain (Smidt and Burbach, 2009), differentiation programs in the developing nervous system are often whereas the Pet1 transcription factor initiates and maintains the expressed in several distinct neuronal cell types, but to what extent terminal differentiation program of serotonergic neurons (Liu et al., they have similar or distinct activities in individual neuronal cell types 2010). However, few neuronal transcription factors are expressed is generally not well explored. We investigate this problem using, as exclusively in only one specific neuronal cell type (Gray et al., 2004; a starting point, the C. elegans LIM homeodomain transcription factor Lein et al., 2007). For example, in addition to being expressed in ttx-3, which acts as a terminal selector to drive the terminal midbrain dopaminergic neurons, Nurr1 is expressed in other non- differentiation program of the cholinergic AIY interneuron class. Using dopaminergic neuronal cell types in which its function is not well a panel of different terminal differentiation markers, including understood, such as the adult olfactory bulb, specific cortical areas neurotransmitter synthesizing enzymes, neurotransmitter receptors and the hippocampus (Zetterström et al., 1996). The expression of a and neuropeptides, we show that ttx-3 also controls the terminal given transcription factor in distinct neuronal populations poses the differentiation program of two additional, distinct neuron types, fundamental question of whether there are underlying common namely the cholinergic AIA interneurons and the serotonergic NSM themes in the activity of the transcription factor in distinct neuronal neurons. We show that the type of differentiation program that is cell types. controlled by ttx-3 in different neuron types is specified by a distinct We have undertaken a systematic, in-depth comparison of the set of collaborating transcription factors. One of the collaborating activity of two transcription factors in the development of several transcription factors is the POU homeobox gene unc-86, which distinct neuronal cell types in the nematode C. elegans, examining collaborates with ttx-3 to determine the identity of the serotonergic whether there are indeed conceptual similarities in the activities of NSM neurons. unc-86 in turn operates independently of ttx-3 in the a given transcription factor in distinct neuron types. We used, as a anterior ganglion where it collaborates with the ARID-type starting point, a member of the LIM homeobox gene family, an transcription factor cfi-1 to determine the cholinergic identity of the ancient family of neuronal patterning genes that display complex IL2 sensory and URA motor neurons. In conclusion, transcription expression patterns in the nervous system of many different species, factors operate as terminal selectors in distinct combinations in from invertebrates to vertebrates (Hobert and Westphal, 2000; different neuron types, defining neuron type-specific identity features. Simmons et al., 2012; Srivastava et al., 2010). One unifying theme is their expression in terminally differentiating neurons (Hobert and KEY WORDS: Caenorhabditis elegans, Homeobox, Neuron Ruvkun, 1998; Moreno et al., 2005). We focus here on the ttx-3 LIM differentiation homeobox gene, which is the sole C. elegans member of the Lhx2/9 subclass of LIM homeobox genes. In vertebrates, Lhx2 is expressed INTRODUCTION in multiple neuronal cell types and is required for the differentiation The development of the nervous system is a multistep process that of olfactory sensory neurons (Hirota and Mombaerts, 2004; employs a series of sequentially acting regulatory factors that Kolterud et al., 2004), the specification of cortical neuron fate successively restrict and determine cellular fates. During the process (Mangale et al., 2008) and the differentiation of thalamic neurons of terminal differentiation, individual neuron types acquire specific, (Peukert et al., 2011). Whether there is a common theme in the hard-wired features that are maintained by the neuron type function of Lhx2 in these distinct neuronal cell types is not known. throughout the life of the animal. A number of transcription factors The C. elegans Lhx2/9 ortholog ttx-3 is exclusively expressed in have been identified that initiate and maintain specific terminal a small number of neurons in distinct head ganglia (Altun-Gultekin differentiation programs in the developing nervous system (Hobert, et al., 2001). ttx-3 null animals display broad differentiation defects 2011). For example, in mouse, the Nurr1 (Nr4a2) transcription in the cholinergic AIY interneuron class. AIY interneurons of ttx-3 null mutants are generated and still express pan-neuronal features, 1Department of Biochemistry and Molecular Biophysics, Columbia University but fail to express scores of terminal identity markers that define the 2 Medical Center, New York, NY 10032, USA. Department of Cell Biology, Yale functional properties of AIY, including genes required to synthesize University School of Medicine, New Haven, CT 06520, USA. 3Instituto de Biomedicina de Valencia, IBV-CSIC, 46010 Valencia, Spain. 4Howard Hughes and package acetylcholine, genes encoding neuropeptide receptors, Medical Institute, Columbia University Medical Center, New York, NY 10032, USA. various types of ion channels and many others (Altun-Gultekin et al., 2001; Hobert et al., 1997; Wenick and Hobert, 2004). TTX-3 *Author for correspondence ([email protected]) exerts this control through direct binding to a cis-regulatory motif Received 31 May 2013; Accepted 11 October 2013 shared by all of its target genes. ttx-3 expression is turned on in the Development 422 RESEARCH ARTICLE Development (2014) doi:10.1242/dev.099721 neuroblast that generates AIY and its expression is maintained One of these factors is the POU homeobox gene unc-86, which is throughout the life of the neuron through an autoregulatory feedback required together with ttx-3 to control the identity of the serotonergic loop (Bertrand and Hobert, 2009) to ensure persistent expression of NSM neurons. unc-86 in turn cooperates with the ARID-type its target genes. A number of transcription factors have been transcription factor cfi-1 to control many terminal identity features described in the C. elegans nervous system that display similar of the cholinergic IL2 sensory and URA motor neurons. Our studies broad-ranging effects on the terminal differentiation programs therefore provide further support for the terminal selector concept executed by the neurons in which they are expressed. These and show that, in combination with other regulatory factors, one transcription factors have been called ‘terminal selectors’ (Hobert, factor can serve as terminal selector in distinct neuronal cell types 2008; Hobert, 2011). It is still an open question how broadly the regulating distinct neuronal differentiation programs. terminal selector concept applies throughout the nervous system; that is, how common it is that many distinct and functionally RESULTS unrelated identity features of a specific neuron type are directly co- Expression pattern of ttx-3 in the C. elegans larval and regulated by a transcription factor or a combination of transcription adult nervous system factors. A ttx-3 reporter gene that contains the ttx-3 locus together with a few Here, we investigate the role of ttx-3 in two additional neuron kilobases upstream but no downstream sequences (ttx-3promA::gfp; classes in which it is normally expressed, namely the cholinergic Fig. 1) was previously shown to be continuously expressed in five AIA interneuron class and the serotonergic NSM neuron class. We distinct neuronal cell types: the cholinergic AIY and AIA find in all three neuron classes that there is a common theme of ttx- interneuron classes, the ASI and ADL chemosensory neuron classes 3 function in that it is broadly required to induce many distinct and and a previously uncharacterized neuronal pair in the pharyngeal functionally unrelated terminal identity features of the respective nervous system (Altun-Gultekin et al., 2001). Transient expression neuron class. Yet the downstream targets of ttx-3 in these neuron was observed in the AIN and SMDD neurons at embryonic stages classes are distinct and are determined by the cooperation of ttx-3 (Bertrand and Hobert, 2009). A fosmid reporter construct, which with a distinct set of transcription factors in different neuron classes. contains more than 30 kb surrounding the ttx-3 locus and which Fig. 1. Expression pattern of the C. elegans ttx-3 LIM homeobox gene. (A) ttx-3 expression constructs and summary of neuronal expression pattern. The promA::gfp and promB::gfp constructs were described previously (Altun-Gultekin et al., 2001; Wenick and Hobert, 2004) and are shown here for comparison only. (B) ttx-3 fosmid expression (wgIs68) in first larval stage animals and in adult animals. D-V, dorsoventral.
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